How the sperm lost its tail: the evolution of aflagellate sperm

The typical sperm is comprised of a head, midpiece and flagellum. Around this theme there is an enormous diversity of form--giant sperm, multi-flagellate sperm and also sperm that lack flagella entirely. Explaining this diversity in sperm morphology is a challenging question that evolutionary biologists have only recently engaged in. Nonetheless, one of the selective forces identified as being an important factor in the evolution of sperm form is sperm competition, which occurs when the sperm of two or more males compete to fertilize a female's ova. In species with a truly monandrous mating system, the absence of sperm competition means that the selection pressure on males to produce motile sperm may be relaxed. Potentially aflagellate sperm are less costly to produce, both in terms of energy and time. Thus, selection may therefore favour the loss of the sperm flagellum and any other motile mechanisms in monandrous taxa. A review of the literature revealed that 36 taxonomic groups, from red algae to fish, were found independently to have evolved aflagellate sperm. I review what is known about the mating systems of each of these taxa and their nearest sister taxa. A sister-group analysis using this information provided weak evidence suggesting that the evolution of aflagellate sperm could be linked to the removal of selective pressures generated by sperm competition.     

The giant sperm of a minute beetle

The mature sperm of Ptinella aptera is a helically coiled, flagellate gamete ca. 1.4 mm long—twice the length of the beetle itself. The rod-like acrosome, comprising the anterior part of the sperm, is 0.4 μm thick but is expanded as a flange around the nucleus and the base of the tail, increasing the diameter of the sperm to 2 μm. The bulk of the tail consists of a pair of bodies with a characteristic ultrastructure of longitudinal tubules beneath a lamellar cortical layer. These bodies are probably homologous with the mitochondrial derivatives of other insect sperm. The axoneme is helically coiled and is flanked by a single accessory body. One of the ‘structured bodies’ is connected to the acrosome and the other to the accessory body. The sperm move actively in the female reproductive tract. The functional significance of this behaviour and of the evolution of the large gamete itself is discussed in relation to existing hypotheses.     

ULTRASTRUCTURE OF THE FLAGELLUM IN SPERM OF CKCINELLA SEPTEMPUNCTATA

Abstract  Using cell whole mount preparation and ultrathin section technique, the ultrastructure of the flagellum in the sperm of Coccinella septempunctata L. was examined with transmission electron microscope. The flagellum is made up of a classic 9+9+2 axoneme containing two similar crystallized mitochondria1 derivatives, two accessory bodies, which are divided in to two portions, an osmiophilic dense crescent and a spongy one, and a non-crystalline body. At the end of the flagellum, only the axoneme is present, it loses the two central microtubules but retains the nine doublets with dynein arms and the nine accessory microtubules.     

A simple kinetic model for dynein oscillatory activity

A kinetic model was proposed for dynein, a motor protein, complexed with microtubule fragments. The model explains the experimental observations of oscillatory movements in surprisingly simple axoneme fragments perfused with an ATP solution. This is the first model explaining the oscillatory activity of dynein as determined by a cooperative interaction of two dynein heads in the axoneme. The oscillation shape, frequency, and amplitude obtained for the model are close to the corresponding parameters determined experimentally.